Comparative population genetics of congeneric limpets across a biogeographic transition zone reveals common patterns of genetic structure and demographic history.

The distribution of genetic diversity is often heterogeneous in space, and it usually correlates with environmental transitions or historical processes that affect demography. The coast of Chile encompasses two biogeographic provinces and spans a broad environmental gradient together with oceanographic processes linked to coastal topography that can affect species' genetic diversity. Here, we evaluated the genetic connectivity and historical demography of four Scurria limpets, S. scurra, S. variabilis, S. ceciliana and S. araucana, between ca. 19° S and 53° S in the Chilean coast using genome-wide SNPs markers. Genetic structure varied among species which was evidenced by species-specific breaks together with two shared breaks. One of the shared breaks was located at 22-25° S and was observed in S. araucana and S. variabilis, while the second break around 31-34° S was shared by three Scurria species. Interestingly, the identified genetic breaks are also shared with other low-disperser invertebrates. Demographic histories show bottlenecks in S. scurra and S. araucana populations and recent population expansion in all species. The shared genetic breaks can be linked to oceanographic features acting as soft barriers to dispersal and also to historical climate, evidencing the utility of comparing multiple and sympatric species to understand the influence of a particular seascape on genetic diversity.

Differential gene expression analysis in the scallop Argopecten purpuratus exposed to altered pH and temperature conditions in an upwelling-influenced farming area.

Increased carbon dioxide in the atmosphere and its absorption across the ocean surface will alter natural variations in pH and temperature levels, occurring in coastal upwelling ecosystems. The scallop Argopecten purpuratus, one of the most economically important species farmed in northern Chile, has been shown to be vulnerable to these environmental drivers. However, the regulatory responses at the gene-level of scallops to these climate stressors remain almost unknown. Consequently, we used an orthogonal experimental design and RNAseq approach to analyze the acute effects of variability in pH and temperature on gene expression in the muscle tissue of A. purpuratus. In respect to control conditions (pH ~ 8.0/ 14 °C), the influence of low pH (~ 7.7) and temperature (14 °C) induced the activation of several genes associated with apoptotic signaling pathways and protein localization to plasma membrane. Elevated temperature (18 °C) and pH (~8.0) conditions increased the expression of transcripts associated with the activation of muscle contraction, regulation, and sarcomere organization effects on muscle tissue. In scallops exposed to low pH and elevated temperature, the genes expressed were differentially associated with the oxidation-reduction process, signal translation, and positive regulation of GTPase activity. These results indicated that the differentially expressed genes under the experimental conditions tested are mainly related to the mitigation of cellular damage and homeostasis control. Our results add knowledge about the function of the adductor muscle in response to stressors in scallops. Furthermore, these results could help in the identification of molecular biomarkers of stress necessary to be integrated into the aquaculture programs for the mitigation of climate change.

Metabolic rate allometry in intertidal mussels across environmental gradients: The role of coastal carbonate system parameters in mediating the effects of latitude and temperature.

We assess the role of direct and indirect effects of coastal environmental drivers (including the parameters of the carbonate system) on energy expenditure (MR) and body mass (M) of the intertidal mussel, Perumytilus purpuratus, across 10 populations distributed over 2800 km along the Southern Eastern Pacific (SEP) coast. We find biogeographic and local variation in carbonate system variables mediates the effects of latitude and temperature on metabolic rate allometry along the SEP coast. Also, the fitted Piecewise Structural Equation models (PSEM) have greater predictive ability (conditional R2 = 0.95) relative to the allometric scaling model (R2 = 0.35). The largest standardized coefficients for MR and M were determined by the influence of temperature and latitude, followed by pCO2, pH, total alkalinity, and salinity. Thus, physiological diversity of P. purpuratus along the SEP coast emerges as the result of direct and indirect effects of biogeographic and local environmental variables.

Population genomic analyses reveal hybridization and marked differences in genetic structure of Scurria limpet sister species with parapatric distributions across the South Eastern Pacific.

The study of sister species that occur in parapatry around biogeographic transition zones can help understand the evolutionary processes that underlie the changes in species composition across biogeographic transition zones. The South Eastern Pacific (SEP) coast is a highly productive coastal system that exhibits a broad biogeographic transition zone around 30-35°S. Here, we present a comparative genome-wide analysis of the sister species Scurria viridula and Scurria zebrina, that occur in parapatry and whose poleward and equatorward range edges intersect in the 30-35°S SEP biogeographic transition zone. We sampled 118 specimens sourced from nine sites from Tocopilla (22°S) to Chiloé (41°S) including one site where both species overlap and analyzed over 8000 biallelic single nucleotide polymorphisms. We found evidence of hybridization between these species in the contact zone and found significant but contrasting population structures for both species. Our results indicate that the genetic structure in S. viridula, which is currently expanding its range poleward, follows a simple isolation by distance model with no traces of natural selection (no evidence of outlier loci). In contrast, S. zebrina, which has  its equatorward range edge at the transition zone, displayed a pronounced genetic break approximately at 32-34°S, along a region of marked environmental heterogeneity in association with a semi-permanent coastal upwelling regime. For S. zebrina we also found 43 outlier loci associated with this genetic break, with a significant proportion of them clustering in a single linkage group. This marked difference in the presence of outlier loci between species suggests that they could be responding differently to local environmental challenges found at their overlapping geographic range edges, thus providing important new insights about genomic changes around biogeographic transition zones in sister species and the forces that shape genetic diversity in intertidal marine species.

Morphological, physiological and behavioral responses of an intertidal snail, Acanthina monodon (Pallas), to projected ocean acidification and cooling water conditions in upwelling ecosystems.

Ocean acidification (OA) is expected to rise towards the end of the 21st century altering the life history traits in marine organisms. Upwelling systems will not escape OA, but unlike other areas of the ocean, cooling effects are expected to intensify in these systems. Regardless, studies evaluating the combined effects of OA and cooling remain scarce. We addressed this gap using a mesocosm system, where we exposed juveniles of the intertidal muricid snail Acanthina monodon to current and projected pCO2 (500 vs. 1500 ppm) and temperature (15 vs. 10 °C) from the southeast Pacific upwelling system. After 9 weeks of experimental exposure to those conditions, we conducted three estimations of growth (wet weight, shell length and shell peristomal length), in addition to measuring calcification, metabolic and feeding rates and the ability of these organisms to return to the normal upright position after being overturned (self-righting). Growth, feeding and calcification rates increased in projected cooling conditions (10 °C) but were unaffected by pCO2 or the interaction between pCO2 and temperature. Instead, metabolic rates were driven by pCO2, but a significant interaction with temperature suggests that in cooler conditions, metabolic rates will increase when associated with high pCO2 levels. Snail self-righting times were not affected across treatments. These results suggest that colder temperatures projected for this area would drive this species growth, feeding and calcification, and consequently, some of its population biology and productivity. However, the snails may need to compensate for the increase in metabolic rates under the effects of ocean acidification. Although A. monodon ability to adjust to individual or combined stressors will likely account for some of the changes described here, our results point to a complex dynamic to take place in intertidal habitats associated with upwelling systems.

Ocean acidification alters anti-predator responses in a competitive dominant intertidal mussel.

Widespread intertidal mussels are exposed to a variety of natural and anthropogenic stressors. Even so, our understanding of the combined influence of stressors such as predation risk and ocean acidification (OA) on these species remains limited. This study examined the response of the purple mussel (Perumytilus purpuratus), a species distributed along Pacific southeastern rocky shores, to the effects of predation risk and OA. Using a laboratory 2 × 2 cross design, purple mussels were either devoid or exposed to predator cues from the muricid snail Acanthina monodon, while simultaneously exposing them to current (500 ppm) or projected OA conditions (1500 ppm). The response of purple mussels to these factors was assessed using growth, calcification, clearance, and metabolic rates, in addition to byssus production. After 60 d, the presence of predator cues reduced mussel growth in width and length, and in the latter case, OA enhanced this response making the effects of predator cues more severe. Calcification rates were driven by the interaction between the two stressors, whereas clearance rates increased only in response to OA, likely explaining some of the growth results. Mussel byssus production also increased with pCO2 but interacted with predation risk: in the absence of predator cues, byssus production increased with OA. These results suggest that projected levels of OA may alter and in some cases prevail over the natural response of purple mussels to predation risk. Considering the role played by this mussel as a dominant competitor and ecosystem engineer in rocky shores, these results have community-wide implications.

Plasticity in organic composition maintains biomechanical performance in shells of juvenile scallops exposed to altered temperature and pH conditions.

The exposure to environmental variations in pH and temperature has proven impacts on benthic ectotherms calcifiers, as evidenced by tradeoffs between physiological processes. However, how these stressors affect structure and functionality of mollusk shells has received less attention. Episodic events of upwelling of deep cold and low pH waters are well documented in eastern boundary systems and may be stressful to mollusks, impairing both physiological and biomechanical performance. These events are projected to become more intense, and extensive in time with ongoing global warming. In this study, we evaluate the independent and interactive effects of temperature and pH on the biomineral and biomechanical properties of Argopecten purpuratus scallop shells. Total organic matter in the shell mineral increased under reduced pH (~ 7.7) and control conditions (pH ~ 8.0). The periostracum layer coating the outer shell surface showed increased protein content under low pH conditions but decreasing sulfate and polysaccharides content. Reduced pH negatively impacts shell density and increases the disorder in the orientation of calcite crystals. At elevated temperatures (18 °C), shell microhardness increased. Other biomechanical properties were not affected by pH/temperature treatments. Thus, under a reduction of 0.3 pH units and low temperature, the response of A. purpuratus was a tradeoff among organic compounds (biopolymer plasticity), density, and crystal organization (mineral plasticity) to maintain shell biomechanical performance, while increased temperature ameliorated the impacts on shell hardness. Biopolymer plasticity was associated with ecophysiological performance, indicating that, under the influence of natural fluctuations in pH and temperature, energetic constraints might be critical in modulating the long-term sustainability of this compensatory mechanism.

Phenotypic plasticity is not a cline: Thermal physiology of an intertidal barnacle over 20° of latitude.

Our understanding of the plastic and evolutionary potential of ectothermic organisms and their populational impacts in the face of rapid global change remains limited. Studies attempting on the relationship between the magnitude of thermal variability across latitude and the degree of phenotypic plasticity exhibited by marine ectotherms are inconclusive. We state that the latter arises from the narrow range of thermal variability captured by the limited span of the latitudinal gradients studied to date. Using a mechanistic ecophysiological approach and a satellite-based assessment of the relevant environmental variables (i.e. temperature and food availability), we studied individuals of the intertidal barnacle Jehlius cirratus from seven local populations widely spread along the Humboldt current system that spanning two biogeographic regions. At the same time, we synthesized published information on the local abundance of our study species across a total of 76 sites representing 20° of latitude, and spanning from 18 to 42°S. We examined the effects of latitude and environmental variability on metabolic rate plasticity, thermal tolerance (thermal breadth and thermal safety margins) and their impacts on the abundance of this widespread marine invertebrate. We demonstrate that the phenotypic plasticity of metabolic rate in J. cirratus populations is not related to latitude. In turn, thermal breadth is explained by the temperature variability each population experiences. Furthermore, we found clinal variation with a poleward decrease of the critical thermal minimum, suggesting that episodic extreme low temperatures represent a ubiquitous selective force on the lower thermal limit for ectotherms. Across our study gradient, plasticity patterns indicate that populations at the equatorial extreme are more vulnerable to a warming climate, while populations located in the biogeographic transitional zone (i.e. high environmental heterogeneity), on the centre of the gradient, display higher levels of phenotypic plasticity and may represent a genetic buffer for the effects of ocean warming. Together, our results suggest the existence of a fitness trade-off involving the metabolic cost of plasticity and population density that is evident only across the vast latitudinal gradient examined.

Nuestro conocimiento del potencial plástico y evolutivo de organismos ectotérmicos y de los posibles impactos poblacionales a la luz del rápido cambio global sigue siendo limitado. Los estudios que relacionan la magnitud de la variabilidad térmica y el grado de plasticidad fenotípica a través de la latitud realizados en organismos ectotérmicos marinos no son concluyentes. Lo anterior creemos que es consecuencia del estrecho rango latitudinal y por consecuencia el menor rango de variabilidad térmica abarcado por los estudios previos. Utilizando un enfoque ecofisiológico mecanicista e información satelital de las variables ambientales relevantes (i.e., temperatura y disponibilidad de alimento), estudiamos individuos del cirripedio intermareal Jehlius cirratus a lo largo de siete poblaciones locales que se distribuyen ampliamente a lo largo del Sistema de la corriente de Humboldt abarcando dos regiones biogeográficas. Al mismo tiempo, sintetizamos la información publicada sobre la abundancia local de nuestro modelo de estudio en un total de 76 sitios que representan 20 grados de latitud y abarcan desde los 18° a los 42°S. Examinamos los efectos de la latitud y la variabilidad ambiental en la plasticidad de la tasa metabólica, la tolerancia térmica (i.e. amplitud térmica y márgenes de seguridad térmica) y los impactos en la abundancia de este invertebrado marino con amplia distribución geográfica. Demostramos que la plasticidad fenotípica de la tasa metabólica en poblaciones de J. cirratus no está relacionada con la latitud. A su vez, la amplitud térmica se explica por la variabilidad térmica que cada población experimenta. Además, encontramos un patrón de variación clinal con una disminución hacia los polos del crítico térmico mínimo, lo que sugiere que las temperaturas episódicas extremadamente bajas representan una fuerza selectiva ubicua en el límite térmico inferior para los ectotermos. A lo largo de nuestro gradiente estudiado, los patrones de plasticidad indican que las poblaciones en el extremo ecuatorial son más vulnerables al calentamiento, mientras que las poblaciones ubicadas en la zona de transición biogeográfica (i.e., alta heterogeneidad ambiental), en el centro del gradiente, muestran mayores niveles de plasticidad fenotípica, lo que puede representar un reservorio genético para los efectos del calentamiento de los océanos. Nuestros resultados sugieren la existencia de un compromiso en la adecuación biológica que involucra el costo metabólico de la plasticidad y la densidad de población que es sólo evidente dado el vasto gradiente latitudinal examinado.

Minimal impact at current environmental concentrations of microplastics on energy balance and physiological rates of the giant mussel Choromytilus chorus.

Microplastic particles (MP) uptake by marine organisms is a phenomenon of global concern. Nevertheless, there is scarce evidence about the impacts of MP on the energy balance of marine invertebrates. We evaluated the mid-term effect of the microplastic ingestion at the current higher environmental concentrations in the ocean on the energy balance of the giant mussel Choromytilus chorus. We exposed juvenile mussels to three concentrations of microplastics (0, 100, and 1000 particles L-1) and evaluated the effect on physiology after 40 days. The impacts of MP on the ecophysiological traits of the mussels were minimum at all the studied concentrations. At intermediate concentrations of MP, Scope for Growth (SFG) had little impact. Other relevant key life-history and physiological processes, such as size and metabolism, were not affected by microplastics. However, individuals treated with MP presented histopathological differences compared to control group, which could result in adverse health effects for mussels.

Biomechanical Characterization of Scallop Shells Exposed to Ocean Acidification and Warming.

Increased carbon dioxide levels (CO2) in the atmosphere triggered a cascade of physical and chemical changes in the ocean surface. Marine organisms producing carbonate shells are regarded as vulnerable to these physical (warming), and chemical (acidification) changes occurring in the oceans. In the last decade, the aquaculture production of the bivalve scallop Argopecten purpuratus (AP) showed declined trends along the Chilean coast. These negative trends have been ascribed to ecophysiological and biomineralization constraints in shell carbonate production. This work experimentally characterizes the biomechanical response of AP scallop shells subjected to climate change scenarios (acidification and warming) via quasi-static tensile and bending tests. The experimental results indicate the adaptation of mechanical properties to hostile growth scenarios in terms of temperature and water acidification. In addition, the mechanical response of the AP subjected to control climate conditions was analyzed with finite element simulations including an anisotropic elastic constitutive model for a two-fold purpose: Firstly, to calibrate the material model parameters using the tensile test curves in two mutually perpendicular directions (representative of the mechanical behavior of the material). Secondly, to validate this characterization procedure in predicting the material's behavior in two mechanical tests.

Upwelling intensity modulates the fitness and physiological performance of coastal species: Implications for the aquaculture of the scallop Argopecten purpuratus in the Humboldt Current System.

Understanding how marine species cope with the natural environmental variability of their native habitats will provide significant information about their sensitivity to the potential environmental changes driven by climate change. In particular, marine species inhabiting upwelling ecosystems are experiencing low seawater temperatures, as well as, acidic and low oxygen conditions as a consequence of the nature of the deep upwelled waters. Our study is focused on one of the most important socio-economical resources of the Humboldt Current System (HCS): the scallop Argopecten purpuratus which has been historically subjected to intensive aquaculture in areas influenced by upwelling processes. Here, a long-term field experiment was performed to understand how tolerant and well-locally-adapted is A. purpuratus to upwelling conditions by studying a set of fitness, physiological, and biomineralogical traits. Stronger upwelling generated a minor water column stratification, with lower temperatures, pH, and oxygen conditions. On the contrary, as upwelling weakened, temperature, pH, and oxygen availability increased. Finally, upwelling intensity also determined the number, duration, and intensity of the cooling and de-oxygenation events occurring in A. purpuratus habitat, as well as, the food availability (chlorophyll-a concentration, Chl-a). Physiologically, A. purpuratus was able to cope with stressful environmental conditions imposed by higher upwelling intensities by enhancing its metabolic and calcification rates, as well, producing higher concentrations of the shell organic matter. These physiological changes impacted the total energy budget, which was highly dependent on Chl-a concentration, and revealed important traits trade-offs with significant fitness costs (higher mortalities emerged when longer and more intense upwelling events succeed). Our study increases the knowledge about the physiological performance and tolerance of this important resource to the ocean acidification and ocean-deoxygenation imposed by variable upwelling intensities, as well as, its potential vulnerability under future changing conditions driven by a potential upwelling intensification.

Discovering divergence in the thermal physiology of intertidal crabs along latitudinal gradients using an integrated approach with machine learning.

In intertidal marine crustaceans, phenotypic variation in physiological and life-history traits is pervasive along latitudinal clines. However, organisms have complex phenotypes, and their traits do not vary independently but rather interact differentially between them, effect that is caused by genetic and/or environmental forces. We evaluated the geographic variation in phenotypic integration of three marine crab species that inhabit different vertical thermal microhabitats of the intertidal zone. We studied seven populations of each species along a latitudinal gradient that spans more than 3000 km of the Chilean coast. Specifically we measured nine physiological traits that are highly related to thermal physiology. Of the nine traits, we selected four that contributed significantly to the observed geographical variation among populations; this variation was then evaluated using mixed linear models and an integrative approach employing machine learning. The results indicate that patterns of physiological variation depend on species vertical microhabitat, which may be subject to chronic or acute environmental variation. The species that inhabit the high- intertidal sites (i.e., exposed to chronic variation) better tolerated thermal stress compared with populations that inhabit the lower intertidal. While those in the low-intertidal only face conditions of acute thermal variation, using to a greater extent the plasticity to face these events. Our main results reflect that (1) species that inhabit the high-intertidal maintain a greater integration between their physiological traits and present lower plasticity than those that inhabit the low-intertidal. (2) Inverse relationship that exists between phenotypic plasticity and phenotypic integration of the physiological traits identified, which could help optimize energy resources. In general, the study of multiple physiological traits provides a more accurate picture of how the thermal traits of organisms vary along temperature gradients especially when exposed to conditions close to tolerance limits.

Transgenerational Effects of pCO2-Driven Ocean Acidification on Adult Mussels Mytilus chilensis Modulate Physiological Response to Multiple Stressors in Larvae.

The effect of CO2-driven ocean acidification (OA) on marine biota has been extensively studied mostly on a single stage of the life cycle. However, the cumulative and population-level response to this global stressor may be biased due to transgenerational effects and their impacts on physiological plasticity. In this study, we exposed adult mussels Mytilus chilensis undergoing gametogenesis to two pCO2 levels (550 and 1200 µatm) for 16 weeks, aiming to understand if prolonged exposure of reproductive individuals to OA can affect the performance of their offspring, which, in turn, were reared under multiple stressors (pCO2, temperature, and dissolved cadmium). Our results indicate dependence between the level of pCO2 of the broodstock (i.e., parental effect) and the performance of larval stages in terms of growth and physiological rates, as a single effect of temperature. While main effects of pCO2 and cadmium were observed for larval growth and ingestion rates, respectively, the combined exposure to stressors had antagonistic effects. Moreover, we found a suppression of feeding activity in post-spawning broodstock upon high pCO2 conditions. Nevertheless, this observation was not reflected in the final weight of the broodstock and oocyte diameter. Due to the ecological and socioeconomic importance of mussels' species around the globe, the potential implications of maternal effects for the physiology, survival, and recruitment of larvae under combined global-change stressors warrant further investigation.

Thermal physiological traits and plasticity of metabolism are sensitive to biogeographic breaks in a rock-pool marine shrimp.

Populations of broadly distributed species commonly exhibit latitudinal variation in thermal tolerance and physiological plasticity. This variation can be interrupted when biogeographic breaks occur across the range of a species, which are known to affect patterns of community structure, abundance and recruitment dynamics. Coastal biogeographic breaks often impose abrupt changes in environmental characteristics driven by oceanographic processes and can affect the physiological responses of populations inhabiting these areas. Here, we examined thermal limits, performances for heart rate and plasticity in metabolic rate of the intertidal shrimp Betaeus emarginatus from seven populations along its latitudinal range (∼3000 km). The distribution of this species encompass two breaks along the southeastern Pacific coast of Chile: the northern break is characterized by sharp discontinuities in upwelling regimes, and the southern break constitutes a major discontinuity in water conditions (temperature, pH, dissolved oxygen and nutrients), coastline topography and divergence of main oceanographic currents. For B. emarginatus, we found higher plasticity in metabolism at the sites sampled at the biogeographic breaks, and at the site subjected to seasonal upwelling. The variation in metabolic rate was not consistent with increasing latitude and it was not affected by breaks. The lower and upper thermal limits were lower in populations around breaks, although the optimum temperature decreased towards higher latitudes. Overall, whereas thermal limits and plasticity of metabolism are related to biogeographic breaks, metabolic rate is not related to increasing latitude or the presence of breaks in the sampled range.

Species-specific responses to ocean acidification should account for local adaptation and adaptive plasticity.

Global stressors, such as ocean acidification, constitute a rapidly emerging and significant problem for marine organisms, ecosystem functioning and services. The coastal ecosystems of the Humboldt Current System (HCS) off Chile harbour a broad physical-chemical latitudinal and temporal gradient with considerable patchiness in local oceanographic conditions. This heterogeneity may, in turn, modulate the specific tolerances of organisms to climate stress in species with populations distributed along this environmental gradient. Negative response ratios are observed in species models (mussels, gastropods and planktonic copepods) exposed to changes in the partial pressure of CO2 (pCO2) far from the average and extreme pCO2 levels experienced in their native habitats. This variability in response between populations reveals the potential role of local adaptation and/or adaptive phenotypic plasticity in increasing resilience of species to environmental change. The growing use of standard ocean acidification scenarios and treatment levels in experimental protocols brings with it a danger that inter-population differences are confounded by the varying environmental conditions naturally experienced by different populations. Here, we propose the use of a simple index taking into account the natural pCO2 variability, for a better interpretation of the potential consequences of ocean acidification on species inhabiting variable coastal ecosystems. Using scenarios that take into account the natural variability will allow understanding of the limits to plasticity across organismal traits, populations and species.

Exploring physiological plasticity and local thermal adaptation in an intertidal crab along a latitudinal cline.

Intertidal organisms have evolved physiological mechanisms that enable them to maintain performance and survive during periods of severe environmental stress with temperatures close to their tolerance limits. The level of these adaptive responses in thermal physiology can vary among populations of broadly distributed species depending on their particular environmental context and genetic backgrounds. Here we examined thermal performances and reaction norms for metabolic rate (MR) and heart rate (HR) of seven populations of the porcelanid crab Petrolisthes violaceus from markedly different thermal environments across the latitudinal gradient of ~3000km. Physiological responses of this intertidal crab under common-garden conditions suggest the absence of local thermal adaptation along the geographic gradient (i.e., lack of latitudinal compensation). Moreover, thermal physiological sensitivities and performances in response to increased temperatures evidenced the existence of some level of: i) metabolic rate control or depression during warm temperature exposures; and ii) homeostasis/canalization (i.e., absence or low levels of plasticity) in physiological traits that may reflect some sort of buffering mechanism in most of the populations. Nevertheless, our results indicate that elevated temperatures can reduce cardiac function but not metabolic rate in high latitude crabs. The lack of congruence between HR and MR supports the idea that energy metabolism in marine invertebrates cannot be inferred from HR and different conclusions regarding geographic differentiation in energy metabolism can be obtained from both physiological traits. Integrating thermal physiology and species range extent can contribute to a better understanding of the likely effects of climate change on natural populations of marine ectotherms.

Biomineralization changes with food supply confer juvenile scallops (Argopecten purpuratus) resistance to ocean acidification.

Future ocean acidification (OA) will affect physiological traits of marine species, with calcifying species being particularly vulnerable. As OA entails high energy demands, particularly during the rapid juvenile growth phase, food supply may play a key role in the response of marine organisms to OA. We experimentally evaluated the role of food supply in modulating physiological responses and biomineralization processes in juveniles of the Chilean scallop, Argopecten purpuratus, that were exposed to control (pH ~ 8.0) and low pH (pH ~ 7.6) conditions using three food supply treatments (high, intermediate, and low). We found that pH and food levels had additive effects on the physiological response of the juvenile scallops. Metabolic rates, shell growth, net calcification, and ingestion rates increased significantly at low pH conditions, independent of food. These physiological responses increased significantly in organisms exposed to intermediate and high levels of food supply. Hence, food supply seems to play a major role modulating organismal response by providing the energetic means to bolster the physiological response of OA stress. On the contrary, the relative expression of chitin synthase, a functional molecule for biomineralization, increased significantly in scallops exposed to low food supply and low pH, which resulted in a thicker periostracum enriched with chitin polysaccharides. Under reduced food and low pH conditions, the adaptive organismal response was to trade-off growth for the expression of biomineralization molecules and altering of the organic composition of shell periostracum, suggesting that the future performance of these calcifiers will depend on the trajectories of both OA and food supply. Thus, incorporating a suite of traits and multiple stressors in future studies of the adaptive organismal response may provide key insights on OA impacts on marine calcifiers.

Geographic variation in thermal physiological performance of the intertidal crab Petrolisthes violaceus along a latitudinal gradient.

Environmental temperature has profound effects on the biological performance and biogeographical distribution of ectothermic species. Variation of this abiotic factor across geographic gradients is expected to produce physiological differentiation and local adaptation of natural populations depending on their thermal tolerances and physiological sensitivities. Here, we studied geographic variation in whole-organism thermal physiology of seven populations of the porcelain crab Petrolisthes violaceus across a latitudinal gradient of 3000 km, characterized by a cline of thermal conditions. Our study found that populations of P. violaceus show no differences in the limits of their thermal performance curves and demonstrate a negative correlation of their optimal temperatures with latitude. Additionally, our findings show that high-latitude populations of P. violaceus exhibit broader thermal tolerances, which is consistent with the climatic variability hypothesis. Interestingly, under a future scenario of warming oceans, the thermal safety margins of P. violaceus indicate that lower latitude populations can physiologically tolerate the ocean-warming scenarios projected by the IPCC for the end of the twenty-first century.

Heritability of hsp70 expression in the beetle Tenebrio molitor: Ontogenetic and environmental effects.

Ectotherms constitute the vast majority of terrestrial biodiversity and are especially likely to be vulnerable to climate warming because their basic physiological functions such as locomotion, growth, and reproduction are strongly influenced by environmental temperature. An integrated view about the effects of global warming will be reached not just establishing how the increase in mean temperature impacts the natural populations but also establishing the effects of the increase in temperature variance. One of the molecular responses that are activated in a cell under a temperature stress is the heat shock protein response (HSP). Some studies that have detected consistent differences among thermal treatments and ontogenetic stages in HSP70 expression have assumed that these differences had a genetic basis and consequently expression would be heritable. We tested for changes in quantitative genetic parameters of HSP70 expression in a half-sib design where individuals of the beetle Tenebrio molitor were maintained in constant and varying thermal environments. We estimated heritability of HSP70 expression using a linear mixed modelling approach in different ontogenetic stages. Expression levels of HSP70 were consistently higher in the variable environment and heritability estimates were low to moderate. The results imply that within each ontogenetic stage additive genetic variance was higher in the variable environment and in adults compared with constant environment and larvae stage, respectively. We found that almost all the genetic correlations across ontogenetic stages and environment were positive. These suggest that directional selection for higher levels of expression in one environment will result in higher expression levels of HSP70 on the other environment for the same ontogenetic stage.

Variation in thermal sensitivity and thermal tolerances in an invasive species across a climatic gradient: lessons from the land snail Cornu aspersum.

The ability of organisms to perform at different temperatures could be described by a continuous nonlinear reaction norm (i.e., thermal performance curve, TPC), in which the phenotypic trait value varies as a function of temperature. Almost any shift in the parameters of this performance curve could highlight the direct effect of temperature on organism fitness, providing a powerful framework for testing thermal adaptation hypotheses. Inter-and intraspecific differences in this performance curve are also reflected in thermal tolerances limits (e.g., critical and lethal limits), influencing the biogeographic patterns of species' distribution. Within this context, here we investigated the intraspecific variation in thermal sensitivities and thermal tolerances in three populations of the invasive snail Cornu aspersum across a geographical gradient, characterized by different climatic conditions. Thus, we examined population differentiation in the TPCs, thermal-coma recovery times, expression of heat-shock proteins and standard metabolic rate (i.e., energetic costs of physiological differentiation). We tested two competing hypotheses regarding thermal adaptation (the "hotter is better" and the generalist-specialist trade-offs). Our results show that the differences in thermal sensitivity among populations of C. aspersum follow a latitudinal pattern, which is likely the result of a combination of thermodynamic constraints ("hotter is better") and thermal adaptations to their local environments (generalist-specialist trade-offs). This finding is also consistent with some thermal tolerance indices such as the Heat-Shock Protein Response and the recovery time from chill-coma. However, mixed responses in the evaluated traits suggest that thermal adaptation in this species is not complete, as we were not able to detect any differences in neither energetic costs of physiological differentiation among populations, nor in the heat-coma recovery.